PHYTOPLANKTON OF TAMPA BAY - A REVIEW

Karen A. Steidinger Florida Department of Natural Resources Bureau of Marine Research 100 Eighth Avenue S.E. St. Petersburg, FL 33701

William E. Gardiner Department of Biology University of South Florida Tampa, FL 33620

INTRODUCTION (Fogg 1965; Loftus et al. 1972; Tundisi Phytoplankton in brackish and 197 1, Herbiand and KuGiller 198 1, and marine habitats, such as estuaries, others). Photosynthetic microorganisms consists of dour principal microalgal less than 20 urn, even Iess than 2 urn, can groups: phytomicroflagellates (7 or more account for 20-90% of the planktonic classes), diatoms, dinoflagellates, and inorganic C uptake, but at the same time blue-green algae. These groups are they can have high respiration rates and representative of several size classes: function in remineralization cycles, thus picoplank ton (0.2 to 2 urn), ul traplank ton being important in detrital food webs (less than 5 urn), nannoplankton (3 to 20 because of their dissolved organic uptake urn or less than 20 urn), and capabilities (Porneroy 1974, Li et g. microplankton (20 to 200 urn). 1983). Number of species and assoxated Occasionally, there are larger species abundance trends in estuaries usually such as Noctituca, which can be 1-2 mm reveal an inverse relationship in'diameter. These algal groups can be horizontally with increasing salinity from found in both the water column and the head to the mouth (Hulburt 1965; sediments of estuaries, yet specific Kinne 1963, and. others). DefeIice and species are usually planktonic or benthic Lynts (1978) also noted an increase in in the prevailing vegetative stage. benthic diatom diversity from terrestriai Benthic microalgal communities (epi- influences to open waters in upper and endolithic, episarnmic, epiphytic) can Florida Bay. Many phytoptankters are be significant components of an estuary cosmopolitan and endemic populations in diversity and biomass. Hustedt (1955) are rare, particularly in estuaries described 89 new species out of 329 (Lackey 1967; Wood 1965; Steidinger species identified in just two mud 1973); however, brackish or marine samples from North Carolina. Dadkr and microaIga1 assemblages (plank tonic and Roessler (1971) gave a biomass figure of benthic) may be distinct areaiiy and 50-630 rng~/rn2for a Florida bay (based seasonally. Most species are "neritic" on chiorophyll conversions) and' Round and represent an estuarinelneritic (1971), in a bent 9.c diatom review, gave grouping although there are periodic 420 mg Chl a/rn . Benthic microalgae invasions of limnetic or oceaniclneritic can serve as food sources for a variety of species due to freshwater discharge or heterotrophs, act as sediment stabilizers, oceanic intrusions respectively (Wood and often become tychoplanktonic in 1965). turbuIent events. Estuaries are dynamic because of Phytoplankton less than 20 urn physical, chemical, and biological often dominate the water column, interactions. The system is a particularly in temperate and subtropical fluctuating, and at times unstable, waters; dominance can be both numerical environment. This affects which species and by biomass, and/or productivity are present as we11 as total production, temporally and spatially. Such a system intensities because of efficient accessory tends to limit planktonic and benthic pigments. populations to those that have wide Temperature has always been used ranges in physioIogica1 and reproductive as a primary factor in distribution and strategies, e.g, most are euryhaline and seasonal occurrence (Braar ud 196 1; eurytherrnal and have "regenerative" or Eppley 1972; Raymont 1980). Many "resting stage" cycles. Additionally, pelagic marine plants and animals are many species have varied nutritional classified geographically by temperature requirements with adaptive assimilation regimes, e.g. boreal, polar, temperate, rates, particularly nannoplankters. and tropical, yet most estuarine Environmental factors limiting phytopiankters appear to be phytoplankton occurrence, diversity, and cosmopolitan. Temperature extremes abundance interact synergistically and and daily differentials can affect d'v principally involve light (water clarity), rates (doublings per day =. 0.8% o. b3?: temperature (metabolic processes, a Q10 of 1-83; Eppley 19721, nutrient division rates), salinity (osmoregulation), availability and uptake, photosynthetic micro- and macronutrients or growth rates, respiration, as imila ion numbers factors, and circulation patterns.' The (mg C mg chl A- I I h the influence of light intensity on primary relationship being An = 1.43ehi ) O.%T at a production is simply demonstrated by the QiO of 2.23 (srnayda 19761, and existence of productivity formulae consequent 1y successional patterns involving solar radiation, extinction because of in terspecific competitive factors, and chlorophyll (Ryther and adaptations. Yentsch 1957; Small g g. 1972; Phytoplankton dynamics, whether Bannister 1974, and others) which are it be in the estuary or in neritic or still used today with some reservations. oceanic waters, have historicaihy been A1 though Ught is required, high levels equated to avaiiabilt y of can cause photoinhi bition and bleaching rnacronutrients. If nitrates, phosphates, of surface restricted organisms. High silicates and the like are in insufficient light intensities can also cause organisms amounts they are considered limiting, to seek a lower level in the water yet growth is a function of cellular column. Diatoms are light saturated at nutrient pools and enzymatic capabilities lower light intensities than and not necessarily external levels dinoflagellates (Riley and Chester 197 11, (Healey 1973). Also, ambient levels of while blue-greens do best at highest light basic nutrients such as NH3-N, N03-N, intensities. Theref ore, adaptations to and PO4-P at specific times do not j light intensity and wavelengths can rates and by reflect turnover cycling j effect competitive advantages. In bacteria, nannopian kters, and animals. addition, photosynthetic organisms have Theoreticaily, macronutrients could be phased uptake and metabolic functions in growth limiting, particularly during both light and dark cycles (see Eppley seasonal bloom periods, with nitrogen 1981). Light also acts to concentrate being considered the. limiting I positively phototactic flagellates in the macronutrient in temperate North euphotic zone so .that there are diurnal Atlantic estuaries (Smayda 1976). or die1 vertical migration patterns However, phytoplankton have varied associated with time of day, light nutritional strategies and adaptations in intensity and water clarity. Benthic relation to availability and uptake, microalgal communities are likewise stored resources, organic N and P affected by light, although the utilization, and growth factor association may be indirect due to requirements, e.g., vitamins, trace substrate preference and its cyclic metals, and chelators. Laws and availability, or circadian rhythms. Most Bannister (I980) stated 'I... phytoplankton photosynthetic benthic microflora can have mechanisms for regulating uptake chromatically adapt to varying light of each element and ... these mechanisms are used to maintain studies, the macrocomponents do not composition and achieve balanced regulate the fine structure of planktonic growth." Even in oiigotrophic oceanic systems. As an example, Provasoli waters, nutrients appear to be (147 1) considered vitamin requirements nonlimiting (see Platt 198 1; Hulburt an important regulatory agent in 1970; McCarthy and Goldman 1979) with occurrence and succession of species. growth rates up to three or more Since the 1960s it has been shown that doublings per day. In constrast, eu- and various phytoplankters, besides bacteria, mesotrophic areas (e.g., Koblentz-Mishke produce vitamins, e.g, BI2, thiamin and and Vedernikos 1976; Redalje and Laws biotin, as well as vitamin inhibitors or 198 1) would appear to have lower growth binders. Recently, Messina and Baker (doublings per day), suggesting (1982). using- axenic cultures of physiological saturation inefficiencies in ~keletbnerna costaturn, Gonyauiax relation to higher nutrient availability tamarensis and Cyclotella cryptica and and "thresh01d'~ growth, or other their - filtrates, showed that & factors. Smayda (1976), in discussing tamarensis and C. cryptica in the Narragansett Bay data, stated exponentiaJ ~rowth..- phase produced high molecular weight - ectdcrines that In 1974, 42% of the annual production inhibited S. costaturn growth responses, of 310 g rn-2 occurred during July and thus effecting a competitive advantage. August when nitrogen remained at The growth response could be very low concentrations. Rapid q'norrnalized'T by addition of excess nutrient recycling was necessary to vitamin or by autociaving the active sustain this production. That it must filtrate prior to testing. Others, have ocurred is consistent with the eating 1978, for freshwater; Freeberg elevated assimilation numbers found --et al. 1979 and Kayser 1981, for marine during the summer despite very low waters) also have used culture filtrates ambient nutrient concentrations ... to demonstrate species succession phytoplankton dynamics during the patterns and that they were mediated by summer in this bay, however, are not competitive inhibition. Provasoii (1979) clearly limited physiologically by pointed out that laboratory experiments nutrient limitation or grazing. While with axenic cultures may produce biased nutrition levels and the recycling rate results because marine bacteria are may set the magnitude (yield) of the capable of enzymatically degrading population pulses, the underlying binders. Provasoli (1979) also discussed factors regulating the fIuctuations in nutrient competition and the importance abundance remain unknown. of chelators, such as hydroxamates, and trace metals (e.g., ferric and cuptic ions) Smayda (1974) had demonstrated earlier in relation to species succession and that bioassays of natural waters would . preconditioning of water masses. Using give more reliable data on limiting -S. costaturn as an example again (it is factors and successional dynamics; he the dominant diatom in temperate and concluded that "...natural phytoplankton tropical estuaries, including Tampa Bay), growth conditioned the surface waters of Morel --et al. (1978) showed that this Narragansett Baym1and that "...chemical species was relativeIy insensitive to changes -in -situ important to the CU"+ activity, yet some other diatoms occurrence, growth, and succession of and ph ytoplankters are sensitive and phytoplankton species would not be- trace metal availability or sequestering detected by the routineiy used nutrient has been used to project species and biological analyses." successions. Such potential species Although there may be correlations interactions are particularly significant between final yield and nitrogen and in estuarine areas with poor flushing, phosphorus levels as suggested by confined water masses, or f rontai Margalei (1971) and shown for some boundaries where biological conditioning and species interactions are accentuated. River (see Lewis and Whitrnan, this Ketchum (19541, among others, volume). Three of these geographic first pointed out the association of subunits connect directly to Lower estuarine circulation with phytoplankton Tampa Bay which opens to the Gulf of accumulation and growth; where there is Mexico through barrier islands. less exchange there can be higher Major freshwater tributaries standing stock because loss due to mixing (Hillsborough, Aiafia, Little Manatee, and export is minimal. Casper (1967) and Manatee Rivers) are located on the also reported that plankton can remain in eastern side and have an average total the same water mass for long periods discharge of about 1350 cfs with peak because of low flushing and mixing - periods usually from June through particularIy in upper reaches of September. Other freshwater drainage estuaries. Later observations and studies enters the bay system through creeks, have emphasized the significance of streams, ditches, canals, ef f iuent pipes, circulation patterns in transport andlor and excessive land runoff. Mean concentration of phytoplankton at a salinities for the system reflect lower density or tidal boundary, motility, and runoff for Old Tampa Bay and the life history stages (e.g., Tyler and Seliger western side of the system with the 1981; Tyler et al. 1982). Most of these greatest influence from the eastern side studies deal with iong estuaries (more due to river discharge. Salinities than I00 miles) and verticalIy stratified increase from an average of 20.8 Oleo at events associated with pycnociines. the u per reaches (Hillsborough Bay) to Wind and tidal action alone can 26.5 8loo mid-bay to 32.2 'loo inside of concentrate -surface organisms by Egmont Key; Gulf salinities just outside accumulation at convergences; with wind the system average 34.1 'loo (McNulty direction and/or intensity changes and --et al. 1972). Because of drainage in tidal changes, patches can be diluted by upper reaches and wind-induced sediment m.ixing and dispersion. Venrick (1978a) suspension in shallow waters, there are stated, "In general, one may expect associated lower water clarity and higher greater phytoplankton heterogeneity in turbidity values which increase or regions of rapid growth and high standing . decrease, respectively, toward the bay crop as well as in regions of mouth. Turbidity also can be due to environmental heterogeneity." Plankton plankton blooms; patches can be on the scale of meters to Mixing of bay waters is tidai or hundreds of kilometers and are wind-induced with a flushing time of influenced by turbulence and other months in upper reaches. The convective processes, growth rates predominately diurnal tides, with versus diffusion, grazing, and community occasional semi-diurnal cycles, create an structure. Platt tal. (1977) suggested average tidai range of 2.3 ft, yet because that patches less than 100 m diameter of natural and man-made constrictions were controlled by turbulence. such as bridges and causeways, circulation and mixing in upper reaches -The Tampa Bay Estuary of the system, i.e. Old Tampa Bay and The Tampa Bay System is a Hiilsborough/McKay Bay, is poor. Poor shallow, vertically mixed, coastal plain circulation and slow mixing contribute to estuary (about 400 mi2) with a mean iong residency times for water masses depth of 3.3 m (see Simon 1974 and this and the biota they support. For this volume for reviews). The system reason, the upper reaches of the estuary consists of eight named subunits: Old often act as nutrient or particulate Tampa Bay, Hillsborough Bay, McKay "sinks." Such conditions can lead to .Bay, Middle and Lower Tampa Bay (more massive algal blooms, anoxia, and the than 50% of the system area), Boca release of H2S. Ciega Bay, Terra Ceia Bay, and Manatee HISTORICAL ACCOUNT OF for the Tampa Bay system. Variables PHYTOPLANKTON STUDIES measured incIuded salinity, temperature, Marshall N., 1952-1953. major nutrients, D.O., pH, turbidity, Marshill (19-e some important chlorophyll, primary production preliminary observations regarding (chiorophyll-light or O2 light-dark phytoplankton standing stock in Tampa bottle), Pt chodiscus brevis (= Bay. "The greatest concentrations, Gymnodinium breve counts,- and limited toward the head of the estuaries studied, phytoplankton identifications and were probably associated with low enumerations. Not all variables were dispersal rates and with mixing of measured at all times for a11 studies; benthic forms into the plankton of most hydrologic collections were shallow water ... the retention of monthly. Their data showed higher phytoplankton populations, permitting standing stock as total chlorophyll 2 in recruitment and growth, may be upper bay reaches with bay system peaks regarded as the more fundamental of in spring and summer. Taylor (1970) these reasons for such contrasts in stated, "The overabundance of production." Although sampling of the phytoplankton in Hillsborough Bay is a bay was not synoptic and represented consequence of high nutrient levels, and only several stations, one or two days is reflected in high figures for during five months in 1952-53, there was chlorophyll 5 and primary production. a north to south trend towards higher Actually, phytoplankters are the only salinities and Iower chIoropi-ryIl a values important primary producers in which is substantiated in later, more Hillsborough Bay because turbidity has comprehensive .studies. reduced light in the littoral zone to such 2. Pomeroy, L. R., 1958. Pomeroy a degree that seagrasses now survive (1960) assessed the "reiativett contribu- only in small, scattered patches." tion of seagrasses and their epiphytes, Dragovich and Kelly (1964; 1966) phytoplankton and benthic microflora in identified limited phytoplankton species Bbca Ciega Bay to primary production during a 1963-64 study. Of the diatoms using the O2 light-dark bottle method or identified, Skele tonema costaturn and changes in dissolved O2 in a bell jar. Asterionella. .- . .. - . . .- - nlacialis were dominant Phytoplankton and benthic microflora species with &- costatum prevalent f rorn were not identified or enumerated. February to May. They also reported Measurements were made on 5 different concur rent dinoflagellate blooms in the days in May and September, 1953 upper system during April 1963: chlorophyll varied from 0.8 to 3.5 mg/m Ceratium hircus (= , furca var. hircus), for water column phytoplankton, and -P. brevis, Prorocentrum micans, and benthic microflora accounted for an Gonyaulax egensis (?I. Dragovich and average of 130 + 30 rng 0 /rn2/hr. He Kelly (19164) reported that concluded that in waters beeper than 2 rnicr'oflagellates of le& than 12 urn m, only phytoplankton were .important dominated Tampa Bay collections during phrnar;<. ~Eobuckrs in Boca Ciega Bay. a portion of 1963. Later, Dragovich-et 3. U.S. Bureau of ~

-*?et a1 this volume. collections showed dramatic weekly (and 4. Florida Board of Conservation daily) dinoflagellate species composition (later the Florida Department of Natural changes. These data, plus offshore data, Resources), St. Petersburg, Florida, 1963 (Steidinger and Williams 1970) to present. These phytoplankton studies demonstrate that monthly phytoplankton were also initiated in response to data collections smooth out or miss major needs for assessing Florida's west coast short-term fiuc tuations and successional red tides and physical, chemical, and patterns. biological correlations. Eldred et i. S teidinger (1973) stated: (1964) reported on the 1963 Tampa Bay red tide and shellfish toxicity in relation ... assemblages that characterize to --P. brevis counts and were the first to eastern Gulf estuarine conditions associate a cause-effect relationship including the diatoms Skeletonema which was later confirmed by McFarren costa turn, Chaetoceros SPP-, (1965) using mouse bioassays. Rhizosolenia stolterf othii, -R. setigera, Saunders et al. (1967) and C ylindrotheca closterium, C yclotelia Steidinger --et aLT967Tstudied diatoms spp., Thalassiosira spp., Thalassionema and dinoflagellates from Tampa Bay to ni tzschioides. Cera taulina ~ela~ica, Caxambas Pass in 1963-1964 at 20 Bacillaria paradoxa, Asterionella- stations; four Tampa Bay stations were japonica, Paralia sulcata, Bacteri- sampled monthly, three were sampled astrurn spp., Nitzschia seriata (?I, weekly and a11 stations but two were less Leptocylindr us danicus, Bellerochea than 2 m deep. Taxa lists included 43 sp., and the dinoflagellates Ceratium diatom genera with 68 identified species, hircus, Gymnodinium splendens, small 21 dinoflagellate genera with 28 Gyrnnodinium and Gyrodinium spp., identified species, 5 blue-green genera Polykrikos spp., Peridinium spp., (e.g. and many unidentified forms. The most P. uin uecorne), Conyaulax spp. (e.g. dominant component of these e.- Torodinium spp., unpreserved samples were micro- Prorocentrum rnicans, and P. gracile flagellates less than 15 urn. Diatorng ... An overview of estuarine, coastal usually were encountered at 1 x 10 (continental shelf) and open Gulf cellsfli ter while dinoflagellates were less phytoplankton reveals four broad types than 20,000/1iter except during blooms. of assemblages: 1) estuarine, 2) ~keletonerna costatim numerically estuarine and coastal, 3) coastal and dominated except during late spring and open Gulf, and 4) open Gulf ... Many summer. In these months, larger species of the diatoms and dinoflage14ates of Rhizosolenia Chae toceros and listed above for estuaries also occur in ~ellerochea(8.horologicalis)dominated coastal waters but usuaIly in lesser the biomass; particularly in lower numbers and lower frequency ... No reaches. El-Sayed --et al. (1972) stated, doubt if we were to carefully identify "The Tampa Bay region has perhaps the the small microflagellates, unarmored most distinct diatom flora of the entire dinoflagellates, and diatoms, we would Gulf. Unusual members of the genera be better able to distinguish an Cydotella, Porosira, Amphipora, estuarine assemblage; however, as Chaetoceros and Bacteriastrum are Saville (1 966) mentioned, estuaries known ... certain species dominate the have few endemic species and are phytoplankton communities regularly ... influenced by freshwater and marine Hemidiscus hardrnanianus- and forms ... Tampa Bay is also rather distinct in that at times it can have Salinities of less than 25 'loo are coincident dinoflagellate blooms, e.g. typically limiting to P. brevis (= G. C rnnodinium breve Con yaulax breve). Steidinger (1975~)also pointed -*th toxic &ecies, one of out that "Gymnodinium breve during coastal origin), Cochlodinium citron, these two outbreaks originaIiy gained Gymnodinium splendens, Gonyautax access to mid-Tampa Bay via the ship spinif era and Peridinium quinquecorne. channel ... once in Tampa Bay ... winds and tides were instrumental in To this list should be added: Ceratium transporting and dispersing blooms." hircus, Prorocentrum micans, P. gracile, Surface counts of P. brevis in Peridinium f oliaceum. ~~hidinium several instances in back-~oca Ciega carterae.. .. Gonvautax diacantha. G. canals were lo7 cells11 due to southwest I---- digitalis, Pyrodmmn baharnense var. winds and incornirrg tides, then on the bahamense, and Noctiluca. Up to five turn of the tide counts were 10~11 species can be co-blooming in the Bay throughout the water column. Prior to system with two to three in the same 1973, researchers believed that red tides general area, particularly the upper started inshore near passes, e.g. off reaches; blooms can be monospecific or Egrnont, and were initiated by previous mixed. Gonyaulax rnonilata typically freshwater drainage, but a reanalysis of blooms in August and September in Old offshore and inshore data suggested red Tampa Bay or mid-bay and is rarely tides started 10-40 miles offshore found during the rest of the year because (Steidinger 1973, 1995% b) and may be the species forms a benthic hypnozygote associated with offshore benthic seed which "overwinters" in the upper 1-5 cm beds in a discrete zone. Offshore of sediment. This species and others initiation has since been demonstrated by that have dormant benthic stages form Haddad and Carder (1979), Steidinger and seed beds; consequently, water column Haddad (1981) and Haddad (19821, and blooms recur in the same area and are has been associated with intrusions of usually seasonal. Recently, Walker and oceanic water on the broad continental Steidinger (197%) were the first to shelf. Walker (1982) recently induce and document the complete documented the sexual cycle of --P. brevis sexual life history of a marine from gamete formation through dinoflagellate - G. moniiata - using planozygotes in culture and is working on Tampa Bay vegzative and zygotic induction of hypnozygotes. 0ther isolates. Steidinger and Ingle (1972) coastal bloom species, e.g. the blue- recounted the I971 Tampa Bay red tide green alga Oscillatoria erythraea, can and Steidinger (19759 stated, aiso be transported into inshore areas and bays. Most monospecific blooms in ... data from two of the three red tides ~arn~a-Bay are dinofla'gellates or blue- that have established in Tampa Bay, greens (e.g. Schizothrix calcicola) that Florida, indicate that the normaily low initiate in the bay itself. salinity barrier of the Tampa Bay Other studies by this agency on System was nonexistent at these times phytoplankton occurring in Tampa Bay because of drought conditions ... include: taxonomic treatments (e.~. saiinites in upper bay reaches, e.g. Old Pyrodinium bahamense, Steidinger --et aj. Tampa Bay, during the summer of 1971 1981; Scrippsiella, S teidinger and Balech were as high as 32°/oo ... high salinity 1977; Gonyauiax, Steidinger 1968, 197 1); conditions in 1971 aiiowed G. breve culture studies (ex. Prorocentrurn -- V blooms, once in the estuary, to minimum (= P. marie-lebourae), Birnhak penetrate and survive in upper reaches and Farrow-1965; Guinardia flaccida, in Old Tampa Bay while salinities in Birnhak --et al. 1967; Ceratulina pelagica, 1974 were at'the lower limit for G. Saunders 1968; Chaetoceros galveston-

breve. densis. Britton and Farrow 1965: Rhodomonas baltica, Detweiler --et al: 1965); documentation of dinoflagellate distribution of chlorophyli -a and its cyst or hypnozygote stages, particularly degradation products in intertidal in bloom species (Walker, unpublished); sediments (Buckley 19821. Of these, and benthic microalgal production of gas Turner% work was the most extensive as bubbles that resuspend and transport regards number of stations (55) and sediment particles. Durako et al. (1982) variables measured, but sampling was demonstrated that dense populations of quarterly from fall 1969 to summer 1971, microalgae, e.g. Peridinium foliaceum thus missing short-term fluctuations and and Gyrodinium fissum in Tampa Bay masking seasonal trends. In addition, the grass flat sediments produced 307.9 ml samples were preserved in formalin, thus of gas/rn2/24 hrs with the highest distorting or lysing most of the production during daylight hours. At unarmored dinoflagellates and other night, there was a 75% reduction in fragile phy toflagellates. Turner bubble production. During the day, measured species occurrence, cell oxygen constituted more than 90% of the counts, biomass, total chlorophy1l 5 gases; at night there was an increase in salinity, temperature, turbidity, depth, C02 composition. These bubbles of water color, total seston, PO4-P, NO3-N, benthic origin were capable of NH3-N, and Si02-Si but found no resuspending sediments by adsorption of consistent significant correlation among warticles. biological and physicochemical variables 5. University of South Florida for most of the collections. Even for the (Department of Marine Science, St. fall 1971 synoptic samples of the bay Petersburg), 1968 to present. Various system (66 samples) Turner stated, "The thesis topics and contract work involving absence of significant correlation among phytoplankton, standing stock, associated factors in the synoptic data of fg11 1971 physicochernical variables, andlor indicates that synoptic sampling may not primary productivity have been reduce many of the problems completed or are underway for the encountered in phytoplankton study." In Tampa Bay area and other nearshore and total, 42 diatom and 23 dinoflagellate coastal areas, e.g. Anclote basin, Crystal taxa were recorded. ~keletonerna River, Charlotte Harbor, and Gulf costaturn was the most abundant diatom waters. These are in addition to in fat1 and spring, particularly in upper unpublished and uncirculated short-term reaches; diatoms' predominated in spring class projects or laboratory reports, e.g. both numerically and in biomass. vertical stratification in Bayboro Harbor Maximum cell counts were recorded in due to runoff and the associated fall with a second maximum in winter. ternporar y separation of freshwater and Summer 1970 had a dinoflagellate marine plankters; the effects of Tampa maximum. Phytoplankton abundance and Bay power plant cooling systems on nutrients generally decreased toward the phytoplankton and standing stock and mouth of the bay. Turner pointed out production; and others. 80th the Platt 5 g.'s (1970) work on spatial Department of Biology and the heterogeneity of phytoplankton where Department of Marine Science use they demonstrated up to a 70% Tampa Bay as a study site. coefficient of variation (CV) for Phytoplankton thesis topics based chlorophyll values between closely on Tampa Bay and offshore Tampa Bay spaced stations (0.5 n mi). The same has have included: description of a new been reported for chlorophyll differences species and distribution of Gonyaulax over one tidal cycle with higher CVs in (Steidinger 197 1); phytoplankton other areas. distribution, abundance and seasonality Turner's recording of oceanic in relation to physicochemical variables Ornithocercus species at salinities of (Turner 1972); oceanic intrusions in 16.7 to 26.9 'loo and counts of up to relation to offshore initiation of red 200,000 ceils/~was in error (~urner,pers. tides (Haddad 19821, and vertical comrn.). The speculation that the Tampa Bay system may be nitrogen Iimited (Gardiner 1982). The latter study is of solely based on N03:P04 ratios (Turner the most interest to this review. and Htlopkins 1974) is beyond the data In a year-long nannopiankton base as outiined in the , Introduction flagellate study, Gardiner (1982) sampled because of unknown & situ turnover five stations in Tampa Bay twice .rates, use of other N sources, such as monthly; one of the stations, Cockroach NH3 and urea, and excess PO4-P values Bay, was sampled for both water and due to natural phosphate beds and mining sediment. Live samples were processed activities (~ohannson 5 , this within 24 hours of collection and relative symposium). abundance of species was estimated. Buckley (1982) reported chlorophyll. At least 25 species were recorded a values for 19 sediment samples from including representatives of the - 2 Lassing Park as 18.3-209.6 mg/m foilowing algal classes: phaeopigments from 13.7-60.9 mg/m , little chlorophyll a was detected below Chlorornonadoph yceae 20 cm. ~hioro~hyila peaked in early Chlorophyceae fall. Since most samp&s were from mud Chrysophyceae and the chiorophyil a values are less than Cryptophyceae some other areas, substrate type, e.g. Dinophyceae firm sand vs. mud, no doubt could Euglenophyceae influence maxima obtained, as could Prasinoph yceae sampling time- in relation to tide and Pr ymnesiophyceae other factors, although Buckley found no distinct vertical migration with This number is conservative, since many sirn uiated tidal cycles. such species superficially resemble More recently, Dr. Gabriel Vargo, a others when examined with light phytoplankton physiological ecologist, microscopy. Accordingly, many of the has been studying N and P metabolism in listed taxa, such as Pyramimonas a Tampa Bay intruder - --P. brevis. His disomata, may include two or more interests are nitrate and urea uptake, morpho1ogicalIy indistinguishable phosphatases, photosynthetic capacity species. Six species were encountered and efficiency, respiration, and vertical year-round including Bi edinomonas migration. He has conducted several rotunda, Cal cornonas laboratory and field trials to date and photosynthetic-k- Chroomonas pluricocca, has verified varied N utilization as well Arnphidinium carterae (in sediments of as a population migration (partial) for Cockroach 13ay),mirnonas disornata,.. .

this species. Eutreptiella marina, Katodiniumy 6. University --of South Florida rotundatum, Gyrodiniurn estuariale and (Department of Biology and Department Prorocen trum minimum. Pr ymnesium of Chemistry), ~arn~&1967 to- present. parvurn is reported to cause fish kills Various theses %and contract work when it reaches densities exceeding involving marine phytoplankton have 100,000 fmt in England coastal waters been completed or are underway. Most (Holdway --et al. 1978). of these studies have concentrated on Other species of interest which toxic dinoflagellates causing red tides were found include the chloromonad and have reported on ultrastructure Hornieiia marina and the chrysophyte (Davis 1969; Steidinger 1979), growth Olisthodiscus iuteus which are requirements (Olander 1968); growth and morphologically similar bloom species.

toxicity (e.g. Doig 1973; McCoy 1977; AH- snecies-1 - identified are between 5 and Eng-W ilmot 1978; Mar tin and Mar tin 20 urn except for Hornielia which is 19761, and ecoiogical impacts (Smith about 50 urn, and larger Amphidinium 1978). One study, unrelated to red tides, carterae.------.- - In- addition to the above work. documented the identity and occurrence two clones of Gyrodiniurn estuariale of phy tomicrof lagellates in Tampa Bay (spring and fall) were tested for growth responses in relation to varied Jight, which agrees with City of Tampa data temperature, and salinity regimes. No for ~chizothrix calcicola. Dino- substantial differences in growth rates flagellates dominated during spring and were detected. summer but were in low concentrations 7. Tampa Electric Company, Big during the rest of the 15 month study; Bend site, (under contract to bloom species were Gonyauiax poly- conservation Consultants, Inc., ramma, G. balechii, G rnnodinium Palmetto, Florida), 1970 to 1977. klsonii (= E.- splendens in Monthly or more frequent (e.g. every 19 - - - . --- days) water samples for chlorophyll, 8. Hiilsborough County Environ- phytoplankton occurrence and mental Protection Commission, (nCEPC) abundance, and certain physicochemical Tampa, 1972 to present. Hillsbo variables were collected from 197'0-1977 county personnel have been sampling the using various field and laboratory Tampa Bay system since 1972; their 50 techniques at several sites in to 54 stations in marine waters were Hillsborough Bay. From 1972-1975, 20 sampled for up to 42 water quality liter samples for phytoplankton were parameters including salinity, concentrated using a 135 urn mesh net; temperature, basic nutrients, chloro- later, a 35 urn net was used. Net phyll, turbidity, dissolved oxygen, light samples were preserved (glutaraldehyde) penetration, and algal blooms. Sampling in one liter; live samples were has been monthly and since 1975, additionally used in 1972-1975. In total, surface, mid-depth, and bottom samples 46 dinoflagellate species and 17 genera, have usually been collected. This data 137 diatom species and varieties and 61 base is available through HCEPC and is genera, and 3 blue-green genera were summarized annually in reports; stored identified (Hughes and Parks 1977). raw data is also available on computer Using the 35 urn data, diatoms had tapes through the Department of primary peaks in winter with secondary Environmental Regulation. Although peaks in late summer/fali. Skeletonema chlorophyll as chlorophyll a is available, costaturn was common and dominant the procedure followed doFs not include with peaks in fall and winter as was grinding and therefore data points may ~hizosoleniasp. cf. delicatula with peaks be underestimated, particularly during in earlv winter. late summer and late diatom peaks or blue-green algal fail. ~keletonehacostaturn abundance blooms. Yet chlorophy11 trends show biased the species diversity index (FIT); north to south decreases. Typically highest diversity occurred at times of upper Old Tampa Bay and Hillsborough/ lowest S. costaturn abundance. This was McKay Bay have chlo ophyll a values also &monstrated by Walker and greater than 20 rng/rn' nd same data S teidinger (1979k) for ~ioridaeast coast points exceed 100 rng/mS during bloom samples and would apply to monospecific events. The lower reach s of the system phytofiagellate bloom events as well. have less than 10 mg/m 5 . The average Other dominants were Herniaulus chlorophyll a concentration for those sinensjs, Oscillatoria sp. (Schizothrix?), stations sampled between 1972-79 was Ceratium hircus, . Asterione Ha glacilis, the hi hest in 1979 with a mean of 18 Gonyauiax polygramrna, Rhizosolenia mg/m B. The lowest for this period was a setigera, Chaetoceros spp., and Nitzschia ' mean of about 10, rng/rn3 in 1972. The spp. This dominance for 15 months is not high chlorophyll a levels for upper necessari t y representative of the system. 1 reaches are often coincident with lower i since bloom events could inf hence salini tylhigher turbidity and lower light i maximum counts and dominance, .and penetration levels. Algal blooms of such events can represent different dinoflagellates, blue-greens and species during- different years. occasionaily diatoms are recorded, but in dsci~~atoria sp. (Schizothrix?) was some instances taxa are oniy identified abundant from late summer through fail to genus or basic taxonomic grouping,

weekly, monthly, quarterly, sporadic) or masses in the upper or middle reaches of analytical techniques (e.g. chlorophyll a the system. Nannoplankters, such as derived with or without grinding of thF small diatoms and microphy toflagellates, concentrated sam pies). Intuitively, these often less than I5 urn, have been specific associations would be suspected reported to dominate water samples because of freshwater inputs, the since 1963. However, not until shallowness of the bay, and poor Gardiner's (1982) dissertation work has circulation and mixing in the upper this group been studied in any reaches where residency of water masses quantitative detail. Prior to Johannson may be months. These trends are (pers. comm.) and Cardiner, few classical associations as expressed by researchers attempted generic or species Hulburt (19651, Casper (19671, Kinne identification because of size and (1~67)~and others. taxonomic difficulties. Dominance of Nanno~lankton(5-20 Nannoplankters not only dominate -urn): As early az965, F~~~ recognized the water column but can be a that at least half of planktonic significant component of benthic photosynthesis was due to microalgae e.g. ~rn~hidiniurncarterae, that passed through a 35 urn mesh net an unarmored dinoflagellate. No one has and that even though the smaller size yet attempted quantitative benthic and classes were the most productive, they epiphytic community studies of went unstudied. Since then there have rnicroalgae in Tampa Bay or other major been numerous studies showing the estuaries in Florida even though biomass importance of pico-, ultra- and as chlorophyll a suggests high standing nannoplankters and their contributions to stocks, and casTal personal observations biomass and primary production in suggest rich communities of diatoms, estuaries, coastal waters, and even open, dinoflagellates and other flagellates. oceanic waters, both in tropical and umber -of Species -and temperate regions. Smaller size classes Ciassification:-- ., . In total. 272 taxa ha;e higher turnover and uptake rates identified to species or varieties have and are metabolically very active. been recorded- from the Tampa Bay Again, Tampa Bay is not an system (see App,endix). If unidentified .. exception to the general rule of forms were added, the list would exceed nannoplankters . dominating micro- 300 taxa. Benthic studies would further planktonic microalgae, except seasonally increase the list and no doubt reveal when monospecific blooms of blue-greens new, undescribed taxa. Of the 272 (~chizothrix) or dinof lageliates - (e.g. specieslvarieties identified, 167 were Gymnodinium nelsonii, Ceratium hircus, diatoms, 78 were dinoflagellates, four Prorocentrum micans, Gonyaulax spp., were blue-greens, and 25 were and others) dominate certain water microphy toflagellates {Table 1).

Table I. Number of Tampa Bay system phytoplankton specieslvarieties and genera by major grouping (see Appendix for species lists and references). I Group Species Genera

diatoms dinof Iagellates blue-greens microflageliates*

~exciusiveof dinoflagellates Comparison of the Tampa Bay system tolerances and growth responses. In at assemblage to other geographic, areas least one species complex shows that 23.5% to 70% of the diatoms (Cr thecodinium cohnii), Beam and from 10 other studies, except two, were Hines-7-r- 1982 identified over 28 sibling also recorded from the Tampa Bay area species and the likelihood of othe; (Tabie 2). The lower the total number of species corn plexes among cosrnopoli tan species recorded, the higher the phy toplankters certainly exists. percentage in common. The two Planktonic versus Benthic exceptions were a North Carolina Assemblages: Planktonic phytoplankton estuary and Florida Bay where benthic have -been well studied world-wide both diatoms dominated. Comparing taxonomically and as primary dinoflagellates from nine of these same producers. However, benthic microalgal studies and another iisting (Table 21, assemblages are not well known even 1 t.3% to 38.1% of the dinoflagellates though they represent a significant were in common. These lower primary production corn onent of an percentages for dinofiageHates, as estimated 100-200 g C/m i'lyr. The lack compared to diatoms, reflect differences of these data from a systems study in temperate versus tropical or approach represents a major gap in our subtropical assemblages. Tampa Bay is knowledge of diversity, function, and strongly influenced by a subtropical energy flow. Diatoms, dinoflagellates, assemblage. In comparing Gulf of and microphy tof lagellates can be epi- Mexico dinoflagellates (El Sayed --et al. and endopelic, episammic, and 1972), 56% of Japanese coastal epiphytic. Some can attach by stalks or dinoflagellates were recorded from Gulf filaments to a variety of substrates; waters whereas only 11.3% were in others live in dose association with common for the Tampa Bay system. The various plant and inorganic materials. I1 phytoplankton studies cited in Tabie 2 Hustedt (1955) identified 319 diatom represent comprehensive studies of species from just two North Carolina eiiher long duration or intensive mud samples; 89 species were new to sampling effort, but techniques differed science. Diatoms, both vegetative and (e.g. live vs. preserved, different sized reproductive stages, can be found down nets) as did sampling frequency and to at least 5 cm as can dinoflageilates, depth. C~nsequentiy, a direct and even down to 10 cm (Round 1971; comparison for similarity indices or Buckley 1982; D. Anderson, Woods Hole association/rnatrix analyses was not Oceanographic Institution, pers. possible nor warranted based on these corn rn.). Vertical distribution is differences and the lack of benthic associated with rhythmic migrations or sampling. biological and physical reworking of Smayda's (1978) biogeographical sediments. Benthic cryptomonads, classif ication scheme based on records chrysomonads, diatoms, and dino- for 379 planktonic species outlined flagellates can be significant components twelve proposed distribution types. of estuarine phytoplankton as pointed out Tampa Bay phytoplankton falls into two by Wood (19651, Saville (19661, Round of his cosmopolitan classes - tern perate (1971) and others. Benthic dino- and warm waters, 'and warm waters. flageilates, such as certain species of Most species are cosmopolitan except for ~r~rocentrurn. Am~hidiniurn.-.- - Theca- those that had been recently described in dinium, Polykrikos, Cymnodinium, the Jast 10 years, e.g. Belierochea Gyrodinium, Scrippsiela. Peridinium, . - horolo~icalis, Conyaulax balechii, and Ostreopsis, and Protoperidinium, can be Proto~eridiniurn steidin~erae.- Even dense and metabolically active in their though many phy toplankters are vegetative stages both in temperate and cosmopolitan in distribution, most tropical areas. For example, Durako et species represent different phys'iologicai -al.9 (1982) study on benthic strains with different environmental dinoflagellate gas production in a Tampa Table 2. Number of species of diatoms and dinoflagellates in common between other geographical areas and the Tampa Bay system. Collection methods, handling methods,numbers of stations, frequency of sampling, and other factors variable.

DIATOMS DINOFLAGELLATES

# % (I%> in B % (#>in Study species Tampa Bay species Tampa Bay Reference

Guadeloupe,West Indies 196 23.5 (46) Ricard & Delesalle 1979 (tropical)

India (tropical) 164 40.0 (66) Subrahrnanyan& Sarrna 1960

Mozarn bique (tropical) 345 29.6 (102) Sournia 1970

Lebanon (war rn temperate) 100 54.0 (54) Lakkis& Novel-Lakkis 1980 0CI NW Atlantic 8 1 63.0 (51) Marshall 1971 (warm tern perateltro pical)

NorthCarolina {temperate) 187 12.8 (24) Campbell 1973

Narragansett Bay Pra tt 1959 (cold temperate)

English Channel (tem,perate) 64 60.9 (39) Maddock et al. 1981

Japan (temperate) 250 36.0 (90) Yamaji 1966

Florida Bay (tropical) 160 13.8 (22) Defelice & Lynts 1978

British Isles Dodge 1981 (cold temperate) Bay seagrass bed suggests high diurnal neritic chain-forming diatom, followed metabolic activity that may indirectly by Rhizosolenia spp., Chaetoceros spp. influence sediment transport by and seasonal dominants such as resuspension via adsorption to bubbles. Bellerochea horologicalis, Schizothrix In addition to benthic vegetative calcicola, and dinoflagellates. cells, this environment harbors a variety Skeietonema numerically dominates the of resting stages of planktonic forms, water column in January to ApriIlMay e.g. hypnozygotes, hypnocysts, and and then again in fall. Bellerochea and resting cells that can influence the Rhizosolenia can dominate in late spring diversity and seasonality of planktonic and summer as can Chaetoceros. diatoms, dinoflageliates and micro- Dinoflagelate blooms, mixed or phytoflagellates. These stages, under monospecific in water masses of long the right conditions, e.g. temperature residency in upper and middle reaches of and photoperiod, act as seed beds for the bay, dominate in summer, fall and future planktonic production, even late spring, depending on particularly in estuaries and coastal environmental factors such as salinity, waters. These "resting" stages have light, temperature, micronutrients, and lowered metabolic acitivity and often organics. These biooms and Schizothrix represent dimorphic life history stages. in upper reaches can lead to oxygen In essence, there are probably few truly depletion in the shaiiow water column holoplanktonic estuarine phytoplankton; and cause fish and invertebrate host are probably meroplanktonic. mortalities. The broad successional Even though benthic phytoplankton pattern, and theref ore seasonality, that can be high in biomass, primary emerges is small diatoms to larger production and species diversity, they diatoms to dinoflagellates to blue- have principally been studied as greens. Margalef (1967) presented a tychoplankton, incidental in water temperate successional pattern of small column coilections. This is an diatoms and flagellates- to Chaetoceros unfortunate oversight. On an areal basis, to Rhizosolenia (both larger diatom primary production of benthic -groups) to coccoiithophorids and microalgae could equal planktonic din~fia~ellates.Even thoLgh there are production values yearly, yet trophically, year to year fluctuations in species benthic microalgae may represent a occurrence and abundance, succession more direct food source to herbivores infers that major seasonal events, e.g. such as ciliates, small crustaceans, and freshwater inputs, temperature, filter or' other suspension feeders than photo period, organic conditioning agents, phytoplankton or detritus from the water and life history strategies kg. benthic column. Not only are there few resting stages), are more important than production values, but turnover rates, macronutrient regimes, at least in intra- and interspecific competition and estuaries. Factors regulating succession, interaction, food webs, morphological for example temperature, affect growth and physiological adaptations, habitat responses and maximum reproductive preference or tolerance, life histories, rate. A species that has a high division diversity, successionat.patterns and other rate (e.g. 2-3 divisions per day or more), factors remain relatively unknown. Such even at the extremes of its temperature studies should be a high priority for tolerance or at a narrow band of high phy toplanktonologists (or phycoiogists temperatures, can ou tcompete another working with microalgae or systems species within a water mass, Such a ecologists if there is a conceptual productive advantage may explain the problem with the term "piankton"). Schizothrix biomass. Blue-greens have Short-term ~luctuations and the highest tolerance to high light and ~uccession]~easonaiity: The most tern perature, followed by dinoflagellates, dominant planktonic species is then diatoms. Smayda's (1974) bioassay Skeletonema costaturn, an estuarine and work suggests that microconstit~ents produced as ectocrines and acting as competitive inhibition or exclusion, or either growth inhibitors or stimulators remain mixed. Those that become regulate succession rather than any monospecific or almost monospecif ic measurable inorganic variable such as often are associated with upper, poorly nitrate, nitrite, ammonia, and ortho- flushed areas and may last months, or at phosphates. It is aIso conceivable that least several weeks. Blooms of Ceratium light, temperature, and salinity influence hircus, Prorocentrum micans, P. gracile, the breaking of dormancy in benthic Gonyauiax digitalis, P tychodiscus brevis,

resting stages and that ectocrines and Schizothrix- - - caicilcola.---- and others have circulation patterns influence dominance lasted several months. Most bloom and abundance of planktonic stages - at species are autochthonous and originate least for dinoflagellates. For example, in the bay system either from a benthic Gonyaulax rnonfiata can bloom ~uly or planktonic inoculum. Two exceptions through September in Tampa Bay and are' --P. brevis and Oscilia toria er ythreae elsewrhere in the Gulf of Mexico; it is which are oceanic/coastal "invaders" and usually absent from the water colbrnn in of allochthonous origin. Blooms of the other months. This species has known toxic dinoflagellate --P. brevis originate seed beds that inoculate the water 10-40 miles offshore, in association with column in Old Tampa Bay. oceanic events, not near passes as Weekly, monthly., quarterly or previously suspected, and can be sporadic sampling misses short-term transported shoreward by currents and fIuctuations that occur daily and winds. Between 1946 and 1982, various tidally. As an example, sampling on an portions of the Tampa Bay system were incoming tide - in back shallow areas exposed to this invader at least 12 times, should reveal a higher composition of usually in the lower reaches, In two eychoplank tonic pennate diatoms. Seven outbreaks, 1963 and 1972, P. brevis to tenfold differences in biomass have penetrated the upper reaches and in one been recorded within one tidal cycle and instance, this species dominated for over this may be due, at least in part, to three months. The only reason P. brevis turbulence and suspension of benthic was able to establish itself in the upper microflora. Sampling at different times portion of the system was because of of the day, regardless of tidal cycle, higher than normal salinity regimes (up may, at least in summer and fall, show to 31 'loo) due to drought conditions at different vertical distribution for the time of the outbreak. In 1963, with f iagellates because of Iigh t adaptations. the lowering of salinities due to rainfall, Night sampling may even reveal the species quickly disappeared, but in migration patterns. Species composition 1971 there was no or minimal rainfall differences occur at "stationstt on a daily during its bloom duration. The species basis during nonbloorn events because a does not survive in waters of less than 25 station is a fixed location while plankton *loo. are transient and water masses are There are several factors rarely tracked. In poorly flushed areas regulating bloom occurrence and of long residency, such differences can duration; some are: seeding of an be negligible. . Because of daily and inoculum, salinity regimes, low mixing of wee kIy differences, seasonality, bay waters, ectocrines, corn peti tive abundance peaks, and successional growth rates, and lack of predators. patterns can be masked by sampling These same factors probably account for infrequency. the Schizothrix blooms, except that the --AUZO~ and Aliochthonous Bloom importance of salinity is replaced by Events: A phytoplankton bloom temperature and light. Again, represents cell densities above monospecific blooms of photosynthetic d&kground levels, usually greater than microalgae, whether the species 50,000 cells/ liter. Such blooms can be produces a toxin or not, can lead to mixed and then monospecific by oxygen depletion and animal mortalities, particularly in early morning hours due example, with the coefficient of to high- respiration activity at night. variation (CV) which does not require a 5arn6ling Strategy -arid ~:al~tical normal distribution of data points around Procedures: Each study should 'have a the mean. sound experimental design prior to Initial sample size and sampling initiation '(see Venrick 1978~-b 5 A). techniques are important to any This planning phase may involve quantitative study because plankton preliminary sampling and analyses to distribution is not homogenous, rather it establish minimal sample size in specific is "patchy" and among sample variation areas or under speciiic conditiohs and is very high (CV greater than 50%). gear type. Sample size can relate to Again, depending on the study goals and volume of field samples, number of field subsequent strategies, different gear replicates, number of subsamples as types that represent continuous replicates, and whether seawater volume sampling, e.g. pumps or nets, may be or cell counts should serve as the sample warranted, or large volume point samples in setting size limitations. When can be pooled and concentrated on analyzing standing stock as chlorophyll board. In reviewing past Tampa Bay or total cell volume, the initial sample system studies, most programs probably size may be in rnl or liters whe had inadequate sampie and subsample unconcentrated, but it may be tens of rn 9 sizes for quantifying abundance or cell if nets are used. When performing volume, and counts should have been diversity or duster analyses, then expressed in orders of magnitude ranges subsample size may relate to cell for relative abundance for among area numbers, e.g. 400 cells per subsample or comparisons using nonpararnetric aliquot with a precision of + 10% of the statistics. mean. Different te&niques or Microscopic examination for approaches may even be used within this identifications and abundance may breakdown. The number of replicates require different volume subsam pies when pooled, or total volume of because, of magnification iimi tations, subsampie, can determine the lower level Many of the srnajler forms have to be of detection if an entire sample is not identified and co,unted under high dry or analyzed, e.g. if ten one rnl replicates (or oil immersion with limited working f 0 mi) are used, the limit of d tec tion is distance, depending on available optics. lo2 celllliter or lo5 cellslrn 5. If only In addition to sample size and one ml is analyz d, the sensitivity i spatial heterogeneity of plankton, fixed decreas d to 109 celis/liter or 10z station data can be misleading; the celtsfm S . Water column samples, in growth medium and the plankton are addition to being just j'trappedW and not transitory. In essence, each collection, concentrated, can be sedimented, even at the same site, represents a new centrifuged, or filtered and the living or area or stations which would preserved concentrate subsampled. theoretically necessitate new sample Again, pooled subsamples or total size determinations. Only in a discrete subsampie size influences the level of water mass which was being tracked or a detection and reported abundance or water mass that was physically retained, biomass. Turner's (1972) level of or even a small lake , would the concept detection was 1 cell = 100,000 of initial sample size have any carryover cellslliter. Lack of adequate sample or meaning for statistical reliablity. subsample size can obviously bias Another probIem with sampling stations numerical data and presence or,absence is whether they should be random or of less abundant forms that may be part source/influence related, e.g. stratified of a characteristic assemblage. The random sampling. For example, in ddvan tage of individual replicates over a Tampa Bay: upper reaches and southern total subsample volume lies in being able boundary areas have freshwater inputs; to determine subsample variation, for water mass residency in Old Tampa Bay and Hillsborough Bay can be 5-27 production, succession, and dynamics, months; shallow areas can contribute the nannopiankton fraction between 5-20 substantial tychoplanktonic corn ponents urn, as we11 as the pico- and from tidal, wind or storm influence, ultraplankton fraction less than 5 urn, etc. Should sampIing be synoptic and at must be studied quantitatively. Also, what depths? There are no simple there are few investigators trained in the answers to llstationtt locations, numer of taxonomy of these small organisms. By stations, or sample and subsample sizes. definition, ecology is based on the study There are other variables suci-r as of biotic and abiotic system components; sampling frequency, gear type, plankton biotic aspects relate to communities and size categories, handling techniques and therefore populations. Populations are strata sampling that also need to be made up of individuals and consequently, addressed depending on project goals and systematics is the grassroots of hypotheses being tested. The questions ecology. It is an indispensable tool that raised above not only apply to unfortunately is looked upon as more of phytoplankton but to zooplankton, an art than a science. Systematic and bacteria, nutrients, turbidity, suspended taxonomic studies are receiving reduced particles, and other aspects of water funding at a time when species column investigations. complexes and an expanded concept of Sampling frequency and timing "speciesw are just being realized. have historically been based on human Most phy toplanktono~ogistsworking and financial resources, sample in the Tampa Bay system and elsewhere processing time, and the convenience of have been preoccupied -with water the investigator. How many column microalgae and the dynamics of planktonologists do night sampling unless planktonic interactions and responses, it is in relation to 24 hour, tidal, or ignoring the microaigat component in the limited cruise sampling? Should benthos. Actually, the two realms are sampling be at a specific tidal phase, e.g. coupled through life histories, vertical high tide, or can sampling be considered migrations and sinking, and yet there are as random when tidal phases are few even qualitative benthic analyses. ignored? Does sampling every second Those benthic studies that are Monday of the month mask daily and quantitative have been selective, weekly fluctuations; does it ignore concentrating on the diatom assern blage, rainfall and runoff events and system the cyst or spore assemblage, or just responses? Quar-terly sampling for biomass as chlorophyll in sediments. certain benthic sessile or inf aunal Such studies cannot be integrated into a populations may be adequate, but plankton dynamics system approach quarterly sampling for water column because of missing compartments. plankton is inadequate unless the Quantitative sampling of benthic investigator is looking at specific within- microalgae, vegetative or reproductive sample correlations and variables in a stages, usually foIlows techniques used large enough data base. for infauna, e.g. replicate core samples; From past and current Tampa Bay sampling could be less frequent than system phytoplankton studies, it is water column collections, but it should obvious that water column nannoplankton take into account rhythms in response to numerically dominate net plankton. Yet tides, light, or other environmental this fraction is usually ignored for its cues. Analyses should be such that living presence and contribution to standing and dead material can be differentiated stock as chlorophyl or primary quantitatively, e.g. by vital staining or productjon. Many of the nannoplankters f tuorescence microscopy, since the are delicate phy tomicrof lagellates that benthos is a detrital reservoir, and do not fix well and therefore are lost to fragile, unpreservable forms should be even baseline data. To understand identified. This necessitates that phytoplankton "diversity", biomass, samples be live and handled within a short time frame to avoid decomposition would total standing stock as active and succession. Cores can be cut for chlorophyll. Recently, Moed and different depths, weighed, sediments Mallegraeff (1978) and Riemann (1978) resuspended in filtered seawater letting have emphasized pH problems with the heavier particles settle out, and spectrophotometric determinations of finally the supernatant size fractioned to phaeopigment concentrations in the remove larger inorganic fractions. This water column. The same problems would procedure would bias the data toward the apply to benthic samples, if not more removed episarnmic assemblage. Or, so. Therefore, it may be best to use sediments from different depths can be chromatographic separation techniques resuspended in filtered seawater of a for benthic samples. known voldme from the site and The above discussion on sampling subsampled with a wide bore pipette. strategy and analytical procedures is Sampling water column phytoplankton superficial at best because it does not with all the associated problems is recommend a final Tampa Bay system technologically easier than sam piing study design for phytoplankton or more benthic populations. Round (197 11, appropriately, microalgae. Such a study Round and Hickman (1971) and Dale would take a team effort of specialists (1 979) gave benthic sampling techniques and a variety of sampling strategies that and strategies for diatoms and shouid address stratified random dinoflagellate cysts and there are several sampling in the water column and accounts for measuring chlorophyll in benthos over a period of years at a sediments. Even presence or absence frequency adequate to characterize without quantitative sampling would be natural fluctuations. Funding and human meaningful data for associations, resources would necessarily restrict seasonality, and Iife history strategies design and scope of such studies and thus since the data gap is so extensive, as limit the usefulness of the data base.

ACKNOWLEDGEMENTS We would like to thank all those that provided unpublished data, and Dr. Gabe Yargo (University of South Florida), Mark Moffler and Michael Durako (Florida Department of Natural Resources) for reviewing the manuscript. Also, we sincerely appreciate the extra time and effort of Virginia Marsh in typing the manuscript.

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DIATOMS (BACILLARIOPH Y cEAE) REFERENCES

Achnanthes longipes Agardh -A. spp. Actinocvclus octdenarius Ehrenber~ (= Actinoptychus *Actinoptychus Ehrenberg (= A. undulatus) -A, spzndens (Shadbolt) Ralf s Amphiprora gigantea var. sulcata (O'Meara) Cleve -A. spp. Amphora marina Van Heurck --A. obtusa Gregory -A. spectabilis Gregory -A. A -A. spp. Anomoeneis s~haero~hora var. scutpta (~hrenber~)Mutler *Asterionella glacialis Castracane (= A. japonica) Asteromphalus flabeljatus (de ~rebisson)Grevilte Auiacodiscus aimberg)A. Schmidt - = Auliscus pruinosus Bailey A. ~unctatusBailev -xi "s-W. ~rnith) Ralf s A. sp. *zaciilaria paxillif er (0.F. ~uller)Hende y (= Nitzchia ~aradoxa) ~acteriastrurn'cdrnosum Pavillard -8. delicatulurn Cleve (= 8. furcaturn ~hadb.sensu ~r~xell) -8. elongaturn CIeve -B. hyalinurn Lauder B. sp. *&llerochea horolo icalis von Stosch (= 8. malleus----I- in part .8idda hia alternans (Bailey) van Heurck &B~I Cleve -3. puichella Gray -D. reticulata Roper REFERENCES -8. tuomeyi (Bailey) Roper - -- Caloneis -westii IW. smith) Hendey Camp1yiodiscus echeneis Ehrenberg {= C. ar~us) "Cerataulina pelameve)Hendey (=-C, bergonii) Chaetoceros affine Lauder

C. affine var. wiliei--- (Grad Hustedt breve Schutt compressurn Lauder crinitum Schutt curv'isetum Cleve danicum Cleve decipiens Cleve didymum Ehrenberg C. diversurn Cleve ?.- m~runow)Meunier ' -C. ~aivestonensisCollier & Mur -C. gracile ScRu tt -C. laciniosurn Schutt -C. lauderi Ralfs -C. lorenzianum Grunow C. pendulum Karsten (ace -C. pelagicum Cleve -C. peruvianurn Brightwell -C.- - sociale Lauder --C. subtile Cleve --C. teres Cleve C. wi hamii Brightwe11 -+-C. 'A' 3 -C. 'B' -C. 'C' -C. 'D' -C. spp. Climacodium biconcavum Cleve -C. f raufeldianum Grunow Cocconeis discuIoides Husted t -C. heteroidea Hantzsch -C. 'A' - -- Corethron criophilum Castracane

*Coscinodiscus centralis- - Ehrenberg -C.-- concinnus- W. Smith -C. curvatulus funo ow -C. granii Gough -C. lineatus Ehrenberg -C. marginatus Ehrenberg -C. radiatus Ehrenberg -C. spp. Coscinosira poi ychorda C ran Cyctotella spp. Dactyliosolen medi terraneus Peragallo -D. sp. REFERENCES

Diploneis crabro Ehrenberg I=Navicula crabro) D. ruendmchmidt) Cleve --T-D. obliqua run) Hustedt -D. 'A' -D. 'B' I3. son. (West) Grunow

. . Epithemia sp. Eucampia cornuta (Cleve) Grunow E. zoodiacus Ehrenber~ - U ~unoto~ramma'marina Eupodiscus argus -E. radiatus Bailey Fragilaria hyalina (Kutzing) Grunow ex Van Huerck F. oceanica Cleve *Erarnmatophora marina {Lyngbye) Kutzing -G. spp. Guinardia flaccida (Castracane) Pergallo G. SDD. - -4t- ~yrosi~mahurnrnii Hustedt -G. wanshec-kin) Cleve Gyrosigma/Pleurosipa 'At Gy rosi~ma/Pieurosigma'B' -G. spp. Hantzschia virgata (Roper) Grunow -H. spp. Haslea wawrikae (Hustedt) Simonsen Herniauius hauckii Grunow -H. rnernbranaceus Cleve H. sinensis Greville *rauderia annulata Cleve (= L. borealis) *Leptocytindrus danicus ~leE -L. miniinus Gran Licmophora abbreviata Agardh .. Li thodesrnium undulatum Ehrenberg -L. spp. Mastigloia binotata (Grunow) Cleve -M. cribrosa Grunow - -. Melosira dubia Kutzing M. ~ranulata~hrenberdRalfs - 1 Agar -M..... 'A'. -M. spp. Navicula cancellata Donkin -N. clavata Gregory -N. f orcipata Greville -N. lyra Ehrenberg -N. marina Ralfs in Pritchard -N. numm ularia ~~ville REFERENCES -N , pelagica -N. pygmea Kutzing -N. 'A' N. spp. *nitzschia closterium (~hrenber~)W Smith -N. constricta Ralfs -N. delica tissima? Cleve (= Pseudonitzschia delicatissirna) N. longissima (de 0rem *K.- pungens var. arlanticum Cleve -N. sigma var. rigida Kutzing -N. 'At -N. '13' -N. spp. Odontella aurita (Lyngbye) Agardh --0. aurita var. obtusa -0. chinensis Grunow (= -0. mobiliensis Grunow --0. obtusa Kutzing (= Biddulphia obtusa)

0. rhombus f. trigona (Cieve -in Van Heurck) - (zmbusf. trigona) *~almerianahardmanianus Greviile (= Hemidiscus hardmanianus) *F?aralia sulcata (Ehrenberg) Cleve Velosirasuicata) -P. sulcata v. coronata (Ehrenberg) Grunow -in Van Heurck -P. 'At Pinnularia rectangulata (Gregory) Cleve -P. 'A' Piagiogramma staurophorum IG regory) Heiberg -P. 'A' -P. spp.-- Plagio~rarnrnopsisvanheurkii (~run~~)Hasle, van Stosch & Svvertsen Pleurosigrna an &turn (Quekett) W. Smith P. balticum (F"1hrenberg W. Smith - -ma balticum) P. fasciola_Y_$_ E~I-=W. Smith - -&ma fasciola) -P. forrnosum W. Smith P. rostoratum Hustedt -P. strigosurn W. Smith -P. 'At -P. spp. Podosira steliiger (~ailey)Mann . Rhabdonema adriaticum Kutzing -R. spg. Rhizosolenia afata Brightwell -R. calcar-avis Schul tze REFERENCES -R. delicatula Cleve -R. fragilissirna Bergon -R. hebatata f. semispina e ens en) Gran -R. imbricata Bri~htwell ~1 robusta.. . ~orrninex Pritchard serigera Brigh tweil stolterf othii Peragallo styliformis Brightwell R. spp. *rkeletonerna costaturn (Greville) Cleve -S. tropicurn Cleve Streptotheca tamesis Shrubsole - Surirella recedens A. Schmidt -S. Rernma(Ehrenberg) Kutzing --S. ovata Kutzing S. spp. Synedra crystallina (Agardh) Kutzing -S. hennedyana Gregory -S. 'At S. spp. *Thalassionema nitzschiodes Grunow -T. spp. Thalassiosira aestivalis Gran & A, ngst *T. decipiens (m~or~ensen *T.- eccenfrica (Ehrenberg) Cleve 1= Coscinodiscus excentricus) T. subtilis bstenfeld) Gran

T. spp. *Thalassiothrix f rauenf eldii (Grunow) Grunow -in Cleve & Gran -T. mediterranea var. pacif ica Cupp T. 'A1 A -T. spp. Trachyneis aspera (Ehrenberg) Cleve -T. sww... Triceratium dubium Brightwell --T. favus Ehrenberrr ~roidoneis lepid&tera (Gregory) CIeve htropislepidoptera)

DINOFLAGELLATES (DINOPH YCEAE)

*Amphidinium carterae Hulburt .A.- crassum Lohmann --A. klebsii Kofoid & Swezy -A. spp. Ceratium furca (Ehrenberg) Claparede & Lachmann REFERENCES

fusus (Ehrenberg) Dujardin hircus Schroder (= --C. furca var. hircus) rnassiiiense (Gourret) Karsten trichoceros (Ehrenberg) Kof oid tripos var. atlanticum (Ostenfeld) Paulsen -C. spp* Cochlodinium citron

I 44 Coolia rnonotis Meunier Dino h sis caudata f. acutiformis Kofoj Skogsber g *-D. cau ata var. peduncuiata (Schmidt) -D. spa ~i~iobsalisIenticula Bergh Fragiiidiurn heterolobatum Balech ex Loeblich

- r. ~on~aulaxbaiechii Steidinger -G. diacanth-nier) SchilIer G. diegensis Kofoid *c,- digitalis (Pouchet) Kofoid C. prindleyi Reinecke 6 Protoceratium reticutaturn *c.-- - monilata Howell- -- -G. polygramma Stein G. scrippsae Kofoid *-.- spinifera (Ciaparede & Lachrnann) Diesing G. 'A' *c mnodiniurn nelsonii Martin -dens in part) ?G.- aurantium Campbell n. sp. -G. spp. & Swezy

C. SOD. -niei (Loebiich) Morrill & Loeblich na .niei).---, ------Katodinium Iwtundaturn {Lohrnann) Loebiich -K*). claucum tebour). Loe blich Nocriluca scintillans (Macartney) Kofoid & Swezy Oblea rotunda ILebour) Baiech ex Sournia T~i~lo~salisrotundurn) Oxyrrhis marina Dujardin Oxytoxum scoinpax Stein 0. SPD. - 1. Proto eridinium abei (Paulsen) 3 *P.+-- claudicans Paulsen) Balech -PI &cum (~;an)Ba~kch P. crassipes (Kofoid) Balech ?F,- deficiens Meunier *P.- depressum (Bailey) Balech -P. divergens (Ehrenberg) Balech -P. excentricurn I~aulsen)Balech -P. nipponicurn be) Balech REFERENCES

*P.- pellucidurn Bergh -P. pentagonurn (Gran) Balech *P. quinquecor~(Abe) Balech P. spiniferum Sckilier) Balech *F. steidin erae Batech I=& oblongurn in part) -+P. striaturn Bohm) " mnis-- (Pavillard) Balech in part) -P. subinerme (Paulsen) Balech --P. tuburnhiller) Balech -P. 'A' -P. SPP. ' ~eridinio~sisas rnrnetrica Mangin Peridinium foliaceum_Y_r Stein) Biecheler -P. aciculif erum Lemmerrnann -P. 'A' -P. 'B' P. 'E' AP. 'N' *Folykrikos hartmannii Zirnmermann P. kofoidii Chatton

*F.A schwartzii Butschlii -P. spp. (0stenfeId) Abe

P. mexicanurn Tafall (=. -P.- maximum in part) *F. micans Ehrenberg *F.- minimum (Pavillard) Sckiller (= & marie-iebouriae in part) -P. oblongurn (~avillard)F. J. R. Taylor P. triestinum Schiller Pt chodiscus brevis (~avis)Steidinger +i=reve) Pyrodinium bahamense var. bahamense Plate P ro hacus horolo 'urn Stein *P-----7-+ stelnu Schiller Wall & Dale P. vancakooae (~ossignol)Wall & Dale ~cri~~siellasubsalsa (6stenfeld) S teidinger & Balech *S. frochoideaLoeblich *Torodiniurn robustum Kofoid & Swezy --T. teredo (P-~ofoid & Swezy Triadinium polyedricurn. .(Pouchet) Dodge (= Goniodoma poi yedricurnl Warnowia SDD...- (=- Pouchetia SDD.) ?Ornithocercus steinii Schutt (Aisidentified) ?O. thumii (~ch~w~ofoid& Skogsberg {mntif ied) NANNOPLANKTON FLAGELLATES REFERENCES {exclusive of Dinophyceae)

CHLOROPHYCEAE

Dunalielia primolecta Butcher

CHRYSOPHYCEAE

Apedinella spinif era (Throndsen) Throndsen Dictyocha fibula Ehrenberg Olisrkodiscus luteus Carter OIisthodiscus carterae Hulburt Mailornonopsis elliptica Matwienko

CRYPTOPHYCEAE

Chroomonas esopus (Conrad & ~uff)Butc

EUGLENOPHYCEAE

Eutreptia lanowii Steur

Eutreatiella marina da-- Cunha- -E. urnnastica Throndsen

Bipedinomonas rotunda Carter Pyramimonas am ylifera Conrad -P. disomata Butcher Tetraselmis apicula ta {Bu tche -T. carteriiforrnis. Butcher -T. xracilis IK ylin) Butcher -T. tetrahele (West) Butcher

PRYMNESIOPHYCEAE

~hrysochrornulina spp. Paviova salina (Carter) Green Pryrnnesium -Carter Hymenomonas carterae (Braarud and Faegerl.) Braarud

CHLOROMONADOPHYCEAE

Horniella marina Subrahrnanyam BLUE GREENS REFERENCES

Agrneneiium spp. (= ~erisrno~edia) Anabaena spp. Anacytis aeruginosa Drouet & Daily Oscillatoria eryth=ea (Ehrenberg) Kutzing Oscilla toria spp. Schizothrix catcicola (Agardh) Comont -S. mexicana Gomont Spirulina spp.

ADDED IN PROOF: Dr. Grethe Hasle (pers. comm.) has identified the following additional diatoms from Tampa Bay: Nitzschia pseudodelicatissima Hasle, Cymatosira belgica Grunow, C. lorenziana Grunow, Skeletonema rnenzeiii Guillard, Carpenter & Reimann, Thalssiosira tenera Proshkina & Laurenko, T. mala Takaro, --T. allenii Takaro, and Minutocellus sp. In addition, Dr, Hasle pointed out the following synonymies: Thalassiosira le to us (=Cascirodiscus lineatus), T. anguste- lineata (=Coscinosira ol%& and Leptocylindrus mexterraneus wyliosolen medi terraneus4